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![Map of distribution of Eudyptula penguins.
Blue and red colours indicate previously-inferred ranges of New Zealand and Australian mitochondrial lineages [18,19], respectively. Lineage distributions in grey dashed areas are unknown. White circles mark sampling localities for the current study. Black lines indicate proximate samples pooled as a priori regional groupings to increase population sample size.](publication/287214614/figure/fig3/AS:339910943887373@1458052493000/Map-of-distribution-of-Eudyptula-penguins-Blue-and-red-colours-indicate.png)
Map of distribution of Eudyptula penguins. Blue and red colours indicate previously-inferred ranges of New Zealand and Australian mitochondrial lineages [18,19], respectively. Lineage distributions in grey dashed areas are unknown. White circles mark sampling localities for the current study. Black lines indicate proximate samples pooled as a priori regional groupings to increase population sample size.
Source publication
Molecular genetic analyses present powerful tools for elucidating demographic and biogeographic histories of taxa. Here we present genetic evidence showing a dynamic history for two cryptic lineages within Eudyptula, the world's smallest penguin. Specifically, we use a suite of genetic markers to reveal that two congeneric taxa ('Australia' and 'Ne...
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Citations
... Eudyptula currently comprises two lineages: the kororāor New Zealand Little Penguin, Eudyptula minor minor Forster, 1781, which is endemic to New Zealand; and the Australian Little Penguin Eudyptula m. novaehollandiae Stephens, 1826. The Australian Little Penguin is native to Australia where the majority of its population occurs, but it also breeds in the Otago region of New Zealand, where it likely arrived after humans first settled in the region (Grosser et al., 2015). The status of these two taxa remains unsettled, with some studies considering them separate species (e.g., Grosser et al., 2017;Cole et al., 2022). ...
A late Pliocene (3.36–3.06 Ma) exposure of the Tangahoe Formation on the North Island of New Zealand preserves close fossil relatives of many extant seabird clades. Here, we report an extinct member of the little penguin ( Eudyptula Bonaparte, 1856) lineage from the Tangahoe Formation—the smallest extinct crown penguin yet known. Eudyptula wilsonae n. sp. is based on the nearly complete skulls of an adult and a fledged but immature individual. Both skulls show more slender proportions than modern little penguins and precede genome-derived estimates for the divergence between Eudyptula minor minor Forster, 1781 (endemic to New Zealand) and Eudyptula m . novaehollandiae Stephens, 1826 (native to Australia and recently established in New Zealand). This raises the possibility that the fossil taxon represents a lineage directly ancestral to extant little penguins. Our results support a Zealandian origin for little penguins, with subsequent Pleistocene dispersal to Australia and a more recent Holocene range expansion of Eudyptula m . novaehollandiae back into New Zealand.
UUID: https://zoobank.org/a415f70a-e441-4920-85e0-75f6457577ea
... The postorbital width in SGO.PV.22245 was estimated as twice the distance between the tip of the processus postorbitalis and the sagittal line because the left side is broken. The systematic classification for modern species followsWinkler et al. (2020) and was updated withGrosser et al. (2015) for Eudyptula species. The stratigraphic scheme concurs with the work of LeRoux et al. (2016), who divided the Bahía Inglesa Formation into eight members. ...
A new cranium of penguin from the Late Miocene of the Bahía Inglesa Formation (Northern Chile) is described here. Specimen SGO.PV.22245 exhibits a unique combination of characters that suggests it belongs to a new species of either Eudypula Bonaparte, 1856, Spheniscus Brisson, 1760, or more probably to a more basal taxon related to them. The specimen is notably smaller than the fossil species of Spheniscus and more similar in size to the extant Spheniscus. The fossa glandulae nasalis narrows caudally, the crista nuchalis transversa and the crista temporalis are expanded like short wings, and a short crista nuchalis sagittalis connects with the rounded and cranially projected eminentia cerebellaris. The fossa temporalis is subtriangular and deeper caudally and the cranial roof is widely expanded. All these features approach the condition to some species of Spheniscus and Eudyptula. However, because of the incompleteness of the material and the fact that it cannot be compared with some fossil species of Spheniscus only known through postcranial material, we are not able to provide a more accurate assignment.
... m. novaehollandiae (Stephens, 1826)) in the south-east of the South Island (within 200 km of Fiordland), and the New Zealand clade (E. m. minor) throughout the country, including the Chatham Islands (Banks et al. 2002;Peucker et al. 2009;Grosser et al. 2015Grosser et al. , 2016Grosser et al. , 2017Checklist Committee 2022). The two clades are sympatric in at least the Otago region, with limited interbreeding, and they are treated as full species by some authors (Grosser et al. 2015(Grosser et al. , 2016(Grosser et al. , 2017. ...
... m. minor) throughout the country, including the Chatham Islands (Banks et al. 2002;Peucker et al. 2009;Grosser et al. 2015Grosser et al. , 2016Grosser et al. , 2017Checklist Committee 2022). The two clades are sympatric in at least the Otago region, with limited interbreeding, and they are treated as full species by some authors (Grosser et al. 2015(Grosser et al. , 2016(Grosser et al. , 2017. Note that subspecies E. m. albosignata, chathamensis, iredalei, and variabilis are no longer recognised, and are treated as synonyms of E. m. minor (Checklist Committee 1990, 2010, 2022, contra Kinsky & Falla 1976, and Checklist Committee 1980. ...
... Note that subspecies E. m. albosignata, chathamensis, iredalei, and variabilis are no longer recognised, and are treated as synonyms of E. m. minor (Checklist Committee 1990, 2010, 2022, contra Kinsky & Falla 1976, and Checklist Committee 1980. The Australian clade of Little Penguin is considered to have colonised New Zealand between AD 1500 and 1900 (Grosser et al. 2015(Grosser et al. , 2016. The western margin of its distribution within New Zealand is poorly known (Grosser et al. 2015) and may have changed over time. ...
... However, the shape of the penguin cornea does vary, with the smaller penguins having smaller, more steeply curved corneas [3]. The most recent transmission (TEM) and scanning (SEM) electron microscopic study of the penguin cornea was performed on what was described as a little penguin Eudyptula minor [4], although we note that these specimens were obtained in Western Australia while ours came from Auckland, New Zealand and recent evidence suggests that they are different species although of the same genus [5][6][7][8][9]. The cornea of both the former penguin and the Magellanic penguin Spheniscus magellanicus have also been the subject of previous electron micrographic studies [2,[10][11][12]. ...
Corneal confocal microscopy has not previously been performed in penguins, despite recognition of its unusually flat shape. To identify features that the penguin shares with other birds and or mammals and those specific to penguins, we undertook confocal microscopic examination of two little (Eudyptula minor), four gentoo (Pygoscelis papua) and five king (Aptenodytes patagonicus) penguin corneas. Transmission electron microscopy was performed on one gentoo and one king penguin, for finer details. Features shared with other higher vertebrates included a five-layered cornea and a similar limbus. Typically avian were a lower density of stromal cells, a more regular arrangement of collagen bands and an absent basal nerve plexus. Features unique to penguins included a flattened superficial epithelium (king penguin), stromal myofibroblasts (all) and an irregular endothelium (little penguin). Other features uniquely identified by confocal microscopy in birds include epithelial and stromal nerves, guttata and stromal imprints on Descemet’s membrane. Transmission electron microscopy identified a lack of wing cells (king penguin), greater posterior collagen lamellae thickness (gentoo penguin) and significantly less interlacing of collagen lamellae in the central cornea (king and gentoo). Most of these unique features are yet to be explained, but some could be adaptations to diving.
... Text added: The Checklist Committee accepts that two Eudyptula taxa are present in New Zealand following the findings of Grosser et al. (2015Grosser et al. ( , 2016Grosser et al. ( , 2017. However, the Committee supports recognition of these taxa as subspecies rather than as species (contra Grosser et al. 2015). ...
... Text added: The Checklist Committee accepts that two Eudyptula taxa are present in New Zealand following the findings of Grosser et al. (2015Grosser et al. ( , 2016Grosser et al. ( , 2017. However, the Committee supports recognition of these taxa as subspecies rather than as species (contra Grosser et al. 2015). Support for these two clades is backed up by earlier results from J. Banks et al. (2008) and Peucker et al. (2009 Widespread throughout coastal North and South Islands and the Chatham Islands but largely replaced by the Australian little penguin in Otago since human settlement (Grosser et al. 2015(Grosser et al. , 2016. ...
... However, the Committee supports recognition of these taxa as subspecies rather than as species (contra Grosser et al. 2015). Support for these two clades is backed up by earlier results from J. Banks et al. (2008) and Peucker et al. (2009 Widespread throughout coastal North and South Islands and the Chatham Islands but largely replaced by the Australian little penguin in Otago since human settlement (Grosser et al. 2015(Grosser et al. , 2016. ...
The fifth edition (2022) of the Checklist of the Birds of New Zealand no longer includes birds from Norfolk Island, Macquarie Island, or the Ross Dependency, Antarctica, unless those species also occur in or have reached New Zealand. Since the publication of the 2010 Checklist of the Birds New Zealand, one previously unknown living taxon (a snipe) has been described, an endemic shag has been split into 2 species, 2 endemic subspecies of petrels have been described, and 11 new vagrant species (3 petrels, 1 booby, 1 shag, 1 ibis, 1 sandpiper, 1 gull, 1 pigeon, and 2 passerines) plus one subspecies (a booby) and two named hybrids (a kiwi and a sandpiper) have been accepted as occurring in New Zealand as at Feb. 2022. The Australian little penguin (Eudyptula minor novaehollandiae) has also been recognised as present and breeding in New Zealand, and the American whimbrel (Numenius hudsonicus) is here recognised as a full species. One vagrant species (black falcon Falco subniger) has been removed from the New Zealand list, crimson rosella (Platycercus elegans) is now considered to be a failed introduction, and the blue shag (= southern populations of the spotted shag Phalacrocorax punctatus) is no longer recognised as a diagnosable taxon. Royal penguin (Eudyptes chrysolophus schlegeli) and Waitaha penguin (Megadyptes antipodes waitaha) are here treated as subspecies rather than full species; and mainland ravens (formerly Corvus antipodum, now Corvus moriorum) are here treated as subspecies of a single species that also occurred on the Chatham Islands, rather than as a full species. The great spotted kiwi (Apteryx maxima) requires this name change, as the type specimens of Apteryx haastii are hybrids between two other species. Eight recently extinct taxa (including two subspecies) have been described or resurrected (2 swans, a duck, 2 penguins, a petrel, a shag, and a parrot), and 30 species that became extinct more than c.
... According to that proposal, the diversity of Eudyptula forms is reflected in the six subspecies inhabiting Australia and New Zealand [2,82]. Other more recent proposals consider that Eudyptula would be constituted by E. minor and E. novaehollandiae, species of recent divergence [83,84] that have been accepted as such by the ornithological community [85]. The inclusion of Eudyptula as the only living species does not modify or bias our results. ...
Despite its current low diversity, the penguin clade (Sphenisciformes) is one of the groups of birds with the most complete fossil record. Likewise, from the evolutionary point of view, it is an interesting group given the adaptations developed for marine life and the extreme climatic occupation capacity that some species have shown. In the present contribution, we reviewed and integrated all of the geographical and phylogenetic information available, together with an exhaustive and updated review of the fossil record, to establish and propose a biogeographic scenario that allows the spatial-temporal reconstruction of the evolutionary history of the Sphenisciformes, discussing our results and those obtained by other authors. This allowed us to understand how some abiotic processes are responsible for the patterns of diversity evidenced both in modern and past lineages. Thus, using the BioGeoBEARS methodology for biogeographic estimation, we were able to reconstruct the biogeographical patterns for the entire group based on the most complete Bayesian phylogeny of the total evidence. As a result, a New Zealand origin for the Sphenisciformes during the late Cretaceous and early Paleocene is indicated, with subsequent dispersal and expansion across Antarctica and southern South America. During the Eocene, there was a remarkable diversification of species and ecological niches in Antarctica, probably associated with the more temperate climatic conditions in the Southern Hemisphere. A wide morphological variability might have developed at the beginning of the Paleogene diversification. During the Oligocene, with the trends towards the freezing of Antarctica and the generalized cooling of the Neogene, there was a turnover that led to the survival (in New Zealand) of the ancestors of the crown Sphenisciform lineages. Later these expanded and diversified across the Southern Hemisphere, strongly linked to the climatic and oceanographic processes of the Miocene. Finally, it should be noted that the Antarctic recolonization and its hostile climatic conditions occurred in some modern lineages during the Pleistocene, possibly due to exaptations that made possible the repeated dispersion through cold waters during the Cenozoic, also allowing the necessary adaptations to live in the tundra during the glaciations.
... Gentoo penguins are broadly distributed around the Southern Ocean and show strong population genetic structuring due to their high residency patterns and foraging behavior close to the coast [35][36][37][38] . The levels of mitochondrial divergence observed are higher than those previously described in rockhopper penguins Eudyptes chrysocome, E. filholi and E. moseleyi 32 and the two putative species of little penguins Eudyptula minor and Eudyptula novaehollandiae 39,40 . In other penguin species, such as chinstrap P. antarcticus 41,42 , macaroni/royal E. chrysolophus and E. schlegeli 32 and king penguin Aptenodytes patagonicus 43 very limited or no population structure is evident throughout their geographical distributions. ...
Although mitochondrial DNA has been widely used in phylogeography, evidence has emerged that factors such as climate, food availability, and environmental pressures that produce high levels of stress can exert a strong influence on mitochondrial genomes, to the point of promoting the persistence of certain genotypes in order to compensate for the metabolic requirements of the local environment. As recently discovered, the gentoo penguins (Pygoscelis papua) comprise four highly divergent lineages across their distribution spanning the Antarctic and sub-Antarctic regions. Gentoo penguins therefore represent a suitable animal model to study adaptive processes across divergent environments. Based on 62 mitogenomes that we obtained from nine locations spanning all four gentoo penguin lineages, we demonstrated lineage-specific nucleotide substitutions for various genes, but only lineage-specific amino acid replacements for the ND1 and ND5 protein-coding genes. Purifying selection (dN/dS < 1) is the main driving force in the protein-coding genes that shape the diversity of mitogenomes in gentoo penguins. Positive selection (dN/dS > 1) was mostly present in codons of the Complex I (NADH genes), supported by two different codon-based methods at the ND1 and ND4 in the most divergent lineages, the eastern gentoo penguin from Crozet and Marion Islands and the southern gentoo penguin from Antarctica respectively. Additionally, ND5 and ATP6 were under selection in the branches of the phylogeny involving all gentoo penguins except the eastern lineage. Our study suggests that local adaptation of gentoo penguins has emerged as a response to environmental variability promoting the fixation of mitochondrial haplotypes in a non-random manner. Mitogenome adaptation is thus likely to have been associated with gentoo penguin diversification across the Southern Ocean and to have promoted their survival in extreme OPEN
... There are several well-documented historic cases of overlumping of species due to the absence of obvious morphological variation among taxa (Adams et al., 2014;Bickford et al., 2007;Grosser et al., 2015). The absence of apparent diagnostic morphological variation in such cryptic or sibling species may be explained by recent diversification, morphological constraints imposed by selective pressures, or by convergence (Fiser et al., 2018). ...
Aim
Delimiting recently diverged species is challenging. During speciation, genetic differentiation may be distributed unevenly across the genome, as different genomic regions can be subject to different selective pressures and evolutionary histories. Reliance on limited numbers of genetic markers that may be underpowered can make species delimitation even more challenging, potentially resulting in taxonomic inconsistencies. Rockhopper penguins of the genus Eudyptes comprise three broadly recognized taxa: northern (E. moseleyi), southern (E. chrysocome) and eastern rockhopper (E. filholi). Their taxonomic status has been controversial for decades, with researchers disagreeing about whether E. chrysocome and E. filholi are distinct species or conspecific. Our goal is to evaluate genome‐wide patterns of divergence to evaluate genetic differentiation and species delimitation in rockhopper penguins, and to assess which mechanisms may underlie previous discordance among nuclear versus mitochondrial analyses.
Location
Sub‐Antarctic and temperate coastal regions of the Southern Hemisphere.
Methods
We generated reduced‐representation genomic libraries using double digest restriction‐site associated DNA (ddRAD) sequencing to evaluate genetic differentiation, contemporary migration rates and admixture among colonies of rockhopper penguins.
Results
The extent of genetic differentiation among the three taxa was consistently higher than population‐level genetic differentiation found within these and other penguin species. There was no evidence of admixture among the three taxa, suggesting the absence of ongoing gene flow among them. Species delimitation analyses based on molecular data, along with other lines of evidence, provide strong support for the taxonomic distinction of three species of rockhopper penguins.
Main conclusions
Our results provide strong support for the existence of three distinct species of rockhopper penguins. The recognition of this taxonomic diversity is crucial for the management and conservation of this widely distributed species group. This study illustrates that widespread dispersive seabird lineages lacking obvious morphological differences may nevertheless have complex evolutionary histories and comprise cryptic species diversity.
... Research has suggested that there are two congeneric taxa of little penguins; Eudyptula novaehollandiae being little penguins occurring in Australia and south east New Zealand (Otago) and Eudyptula minor being little penguins occurring in all other parts of New Zealand (Peucker et al. 2009;Grosser et al. 2015Grosser et al. , 2016Grosser et al. , 2017. This genetic difference has been suggested to be linked to some morphological and behavioural differences between these two distributions of little penguins (Grosser et al. 2015(Grosser et al. , 2017. ...
... Research has suggested that there are two congeneric taxa of little penguins; Eudyptula novaehollandiae being little penguins occurring in Australia and south east New Zealand (Otago) and Eudyptula minor being little penguins occurring in all other parts of New Zealand (Peucker et al. 2009;Grosser et al. 2015Grosser et al. , 2016Grosser et al. , 2017. This genetic difference has been suggested to be linked to some morphological and behavioural differences between these two distributions of little penguins (Grosser et al. 2015(Grosser et al. , 2017. For example, it has been reported that Australian little penguins commonly come ashore after dusk in 'rafts' where they swim ashore in groups, walk together to their nest areas and are capable of double brooding (Grosser et al. 2015). ...
... This genetic difference has been suggested to be linked to some morphological and behavioural differences between these two distributions of little penguins (Grosser et al. 2015(Grosser et al. , 2017. For example, it has been reported that Australian little penguins commonly come ashore after dusk in 'rafts' where they swim ashore in groups, walk together to their nest areas and are capable of double brooding (Grosser et al. 2015). These behaviours have not been as frequently observed in the New Zealand lineage of little penguins potentially because of genetic differences. ...
Little penguins (Eudyptula minor), or Kororā in Māori, show variation in their behavioural responses towards zoo visitors in Australian zoos. We experimentally examined the effects of visitor presence on the behaviour and stress physiology of little penguins at Wellington Zoo, New Zealand. The two treatments were: (1) Visitor presence – the exhibit was open to visitors; and (2) Visitor absence – the exhibit was closed to visitors. We found that when the exhibit was closed to visitors, the percentage of penguins observed close to the visitor viewing pier increased from about 1% to 9%, but there was little effect on other behaviours and faecal glucocorticoid metabolite concentrations. This result of increased avoidance of the visitor viewing pier when the exhibit was open to visitors suggests close visitor contact, particularly from above, may be fear-provoking for these penguins. We conclude that designing enclosures to allow close viewing proximity of visitors, such as visitors looming over the pool, may be futile in improving visitor experience, since this species of penguins is likely to avoid these types of viewing areas when visitors are present. Visitors positioned close to and above penguins may be particularly problematic since visitors in this position may be perceived as threatening.
... A detailed understanding of the genetic structure of declining populations is essential to develop appropriate conservation actions (e.g., Ben-Menni Schuler et al. 2017;Forrest et al. 2017;Moura et al. 2019). We agree with Burridge (2020) that little penguins (Eudyptula minor; also referred as Eudyptula novaehollandiae: Grosser et al. 2015) are a challenge for conventional conservation genetic analyses. The commentary by Burridge (2020) raised 4 main concerns: 1) that the selection of genetic clusters by Colombelli-Négrel et al. (2020) was contrary to the long-standing approach published by Pritchard et al. (2010) and adopted by many genetic studies; 2) that the gene flow results reported by Colombelli-Négrel et al. (2020) should be discounted; 3) that management recommendations for a single population should not be made based on genetic clustering; and 4) that Colombelli-Négrel et al. (2020) falsely represented and misinterpreted published related work in little penguins. ...
